Forest communities with Hacquetia epipactis (Scop.) DC. in the Cieszyńskie Foothills. 1. Introduction and aim of the study. original research article

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1 DOI: /frp Available online: original research article Leśne Prace Badawcze / Forest Research Papers September 2015, Vol. 76 (3): e-issn Forest communities with Hacquetia epipactis (Scop.) DC. in the Cieszyńskie Foothills Magdalena Mijal Forest Disctrict Wisła, ul. Czarne 6, Wisła, Poland Tel , magdamijal@interia.pl Abstract. The strictly protected Hacquetia epipactis, which natural range has its north-eastern limit in the area of the Cieszyńskie Foothills, is one of the most interesting rarities of Polish flora. For one and a half centuries H. epipactis was popular with scientists resulting in a relatively high number of papers addressing this plant. Usually, however, the characteristics of sites where H. epipactis occurs have been overlooked by the researchers. The aim of this work was to fill this gap, at least partly, by determining plant associations that are preferred by this species. Based on previous phytosociological records and by implementing numerical classification methods, it was found that the Hacquetio-Fraxinetum excelsioris Marinček 1990, which is divided into two subassociations (typicum and allietosum with other variants), is the main association in which this plant species occurs. In isolated sites, mostly outside the Cieszyńskie Foothills, it also appears in fragments of Galio sylvatici-carpinetum betuli Oberd and Tilio cordatae-carpinetum betuli Tracz associations. Keywords: endangered species, plant communities, Hacquetio-Fraxinetum excelsioris, phytosociology, rare plant association 1. Introduction and aim of the study The north-eastern edge of the territorial range of Hacquetia epipactis (Scop.) DC of the Umbelliferae (Apiaceae) family is found in Poland. Its sites in the Silesian areas of and Opole were the first ones described in today s Poland (Wimmer 1857), although the plant presumably was known in this area much earlier. Evidence of this observation is found, among others, in the specimens of a herbarium collected in 1830, which today is part of the herbarium collection of the Museum of Natural History at the University of Wrocław (Guzikowa 1970; Duda et al. 2001; Henel 2006). Many researchers have devoted their attention to Hacquetia epipactis, mainly in determining its distribution in Europe, but also in describing its biology, morphology, ecology, classification of plant communities, protection, and even restoration, pharmacology and ethnography (Kolbenheyer 1862; Hegi 1926; Szafer 1929; Kozłowska 1936; Tumidajowicz 1964; Guzikowa 1970; Marinček 1990; Karcz, Trząski 1995; Duda et al. 2001; Henel 2006; Gajewski et al. 2011; Wika et al. 2014). The popularity of Hacquetia epipactis among the inhabitants of the Cieszyńskie Foothills did not always benefit the plant, which was already noted at the beginning of the 20th century (Simm 1924). Frequent flower picking and plant removal contributed to the reduction of its population (Guzik et al. 2008). Hacquetia was listed as a fully protected species only on 9 July, as a result of the effort of naturalists. It is also on the Red list of plants and fungi in Poland as a species vulnerable to extinction in isolated stands outside of its main area of occurrence (Zarzycki, Szeląg 2006). Research on the classification of plant communities which include Hacquetia epipactis in Poland, including the Cieszyńskie Foothills, as well as other parts of Europe, are incomplete. The adjoining Moravian Gate indicates a connection between the Polish sites of H. epipactis with sites in the southern part of its range. Therefore, it is reasonable to develop a classification of plant communities to protect the specific habitats and plant communities where H. epipactis is found. Despite many studies about this plant, no full phytosociological characterization exists or if so, it is limited to a specific, isolated site. The aim of this paper is to characterize the phytocoenoses of Hacquetia epipactis and to classify the plant communities where it occurs on the basis of collected phytosociological documentation. Submitted: , reviewed: , M. Mijal

2 274 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Methodology The study included most of the known and new sites of Hacquetia epipactis in the Cieszyńskie Foothills (Pelc 1969), as well as some sites outside of this area (Tumidajowicz 1964; Guzikowa 1970; Zając, Zając 2001; Henel 2006). To carry out the analysis, 46 phytosociological records were made in selected areas using the Braun-Blanquet method (Pawłowski 1972) and 36 phytosociological records were used from the archives of the Department of Forest Botany and Nature Protection of the Agricultural Academy of Kraków (currently the Department of Forestry Biodiversity, Institute of Ecology and Silviculture, University of Agriculture, Kraków) (Skrzydłowski 2000; Różański 2001a, 2001b). The numerical classification was performed for a total of 82 phytosociological records, placed on an ATPOL grid (Fig.1). The recorded sites of Hacquetia epipactis were located at an altitude of meters above sea level. Cluster analysis was used to determine the relationship of the similarity between each pair of compared phytosociological records (Sneath, Sokal 1973; Dzwonko 1977a, 1977b). Due to the non-vectorial nature of the Braun Blanquet scale, the results had to be transformed into numerical data. We adopted a method in which species of low quantity have a relatively significant impact on the similarity between the compared records. In order to calculate the similarity between phytosociological records for qualitative and quantitative data, the modified Marczewski Steinhaus index was used (Różański 1988). The matrix of similarities between each pair of phytosociological records was classified according to the agglomeration of unweighted pair group method with arithmetic mean (UPGMA) (Sneath, Sokal 1973 Dzwonko 1977a, 1977b). As a result, grouping the phytosociological records by cluster analysis produced dendrograms of qualitative and quantitative data. A dispersion diagram is produced when they are assembled together (Dzwonko, Grodzińska Figure 1. Localities of phytosociological records with Hacquetia epipactis in ATPOL grid system. Explanations: PC Cieszyńskie Foothills with one plot in the Śląskie Foothills (26), GR Grudzice, SO Opole Silesia, OGW Upper Warta Depression, MO Mogilany, KZ Żywiec Valley, BS Silesian Beskid.

3 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): ). The groups distinguished in this manner were used for the final classification of the analysed records and phytosociological tables were produced using the classical methods of observation and comparison (Pawłowski 1972). The study includes only those phytosociological tables for the distinguished Hacquetio-Fraxinetum excelsioris communities. The calculations and figures in this part of the study were performed using Canoco and Statistica statistical software. The nomenclature of vascular plants is from Mirek et al. (2002). 3. Results Classification of the plant communities with Hacquetia epipactis In order to classify particular types of forest communities selected for the presence of one plant, a dendrogram pair was used, calculated for qualitative and quantitative data (Fig. 2, 3). A record as included in the same community when its similarity was no less than 0.33, although depending on the needs of study, this criterion can be somewhat modified (Dzwonko 1977a). The analysis of all 82 records in the dispersion diagram (Fig. 4) ultimately resulted in the identification of 74 records, with the remaining eight unclassified. Based on a qualitative scale, two large groups were identified as sub-associations: typicum (Table 1) and allietosum (Table 2) of the Hacquetio-Fraxinetum excelsioris association, as well as two small groups, designated as fragments of the Tilio cordatae-carpinetum betuli and Galio sylvatici-carpinetum betuli associations. On the other hand, based on quantitative scale five sub-association s variants were identified. The final classification of the distinguished plant, confirming the results of the studies of Różański (2001b) and Różański and Dziedzic (2003), according to the taxonomy of Polish communities (Matuszkiewicz 2005) is as follows: Class: Querco-Fagetea Br.-Bl. et Vlieg. 1937; Order: Fagetalia sylvaticae Pawł. in Pawł. Sokoł. et Wall. 1928; Alliance: Tilio platyphyllis-acerion pseudoplatani Klika 1955; Association: Hacquetio-Fraxinetum excelsioris Marinček 1990; Sub-association: Hacquetio-Fraxinetum excelsioris allietosum 34 records; Variants: Typical 22 records; With Fagus sylvatica 12 records; Sub-association: Hacquetio-Fraxinetum excelsioris typicum 30 records; Variants: With Ulmus scabra 12 records; Figure 2. Dendrogram of phytosociological records similarity made for Hacquetia epipactis plots on Cieszyńskie Foothills and in the selected places in Poland according to quantitative scale Poor 9 records; Disturbed 9 records; Alliance: Carpinion betuli Issl em. Oberd. 1953; Association: Tilio cordatae-carpinetum betuli Tracz records; Association: Galio sylvatici-carpinetum betuli Oberd records.

4 276 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Figure 3. Dendrogram of phytosociological records similarity made for Hacquetia epipactis plots on Cieszyńskie Foothills and in the selected places in Poland according to transformated quantitative scale Characteristics of forests with Hacquetia epipactis 1. Tilio platyphyllis-acerion pseudoplatani alliance 1.1. Hacquetio-Fraxinetum excelsioris association The vast majority of forests or their fragments in the Cieszyńskie Foothills part of this association, whose characteristic plant is primarily Hacquetia epipactis, as well as Fragaria moschata, Veronica montana, Actaea spicata, Orchis pallens, Thalictrum aquilegifolium and locally Arum alpinum. Distinguishing this new association in Poland has sorted out previously existing inconsistencies in the classification of different, multi-species, fertile deciduous forests in the Cieszyńskie Foothills. In numerically analyzing the records, two different sub-associations of this association were distinguished typicum and allietosum, which were further divided into variants (Fig. 4) Sub-association typical of Hacquetio-Fraxinetum excelsioris typicum (Table 1) The main distinguishing feature of this sub-association is the almost complete lack of Allium ursinum. The frequency and coverage of the characteristic and differential species of the association, including Hacquetia epipactis, are much greater than in the allietosum sub-association. Fraxinus excelsior and Quercus robur form the tree layer, with Tilia cordata, Acer pseudoplatanus and Ulmus scabra also occurring. The artificially introduced species of Picea abies and Larix decidua are also found here, while Fagus sylvatica and Acer campestre are very rare. The second tree layer most often consists of Carpinus betulus, which is also present in the upper layer, with Tilia cordata and Acer platanoides in some patches. The shrub layer in this sub -association is poor, with only Sambucus nigra, Lonicera xylosteum and Corylus avellana occurring in significant amounts. The forest floor is covered in the range of 55 85% and there is no single dominant species. Some of the phytocoenoses are distinguished by a poor forest floor or tree layer. They are characterised by a significant share of Anemone nemorosa, Mercurialis perennis, Galeobdolon luteum, Symphytum tuberosum, and Ficaria verna. Hacquetia epipactis found in this sub-association has optimal conditions for development, forming large clumps with short stems, and is not threatened by competition. Three variants were distinguished in the typicum sub-association: (1) with Ulmus scabra, (2) poor and (3) disturbed: The Ulmus scabra variant seems to represent the most typical form of the association, even though it is not found in as fertile habitats as the allietosum sub-association. The tree stand almost always has two layers and is generally well preserved. The upper, emergent layer consists of Fraxinus excelsior and Quercus robur, but also Ulmus scabra, Tilia cordata, and less frequently Acer pseudoplatanus and A. platanoides. The second, tree layer is formed mostly of Tilia cordata, Carpinus betulus as well as Acer platanoides and Ulmus scabra. The shrub layer is not very varied, consisting of Sambucus nigra, a few Crataegus monogyna and Crataegus laevigata as well as Corylus avellana. The herb layer has relatively large numbers of Hacquetia epipactis, with an average coverage of 70%. Additional species most commonly include: Galeobdolon luteum, Asarum europaeum, Symphytum tuberosum, as well as Anemone nemorosa and Ficaria verna. In addition to Hacquetia epipactis, one of the characteristic Hacquetio-Fraxinetum excelsioris association species also found here is the locally

5 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Figure 4. Dispersion diagram made of two dendrograms according to quality and quantity scale for 82 phytosociological records with Hacquetia epipactis with determined syntaxa. Classification of plant communities: I association Hacquetio-Fraxinetum excelsioris: I.I sub-association Hacquetio-Fraxinetum excelsioris allietosum, A typical variant, B Fagus sylvatica variant, I.II sub-association Hacquetio-Fraxinetum excelsioris typicum, C Ulmus scabra variant, D poor variant (vegetation quantity) E disturbed variant, II Carpinion betuli: F Tilio cordatae-carpinetum betuli, G Galio sylvatici-carpinetum betuli,without frame with letter unclassified phytosociological records differential Arum alpinum. No species from the Vaccinio- Pice-etea class were noted; Carpinion betuli species and other species were also rarely found. Species of the Fagetalia sylvaticae order and the Querco-Fagetea class dominated The poor variant is primarily distinguished by the clearly distorted layer of trees. Artificially planted Quercus robur and Picea abies often dominate here. Gaps in the upper tree layer are filled by Carpinus betulus and Acer pseudoplatanus. The tree stand is actually characterised by a single-layered structure. The shrub layer consists of large numbers of only Lonicera xylosteum and Sambucus nigra. No species characteristic for the association is found in the herb layer, except for the small number of Hacquetia epipactis. Only the Fagetalia sylvaticae order is represented by a large number of characteristic species, in even greater numbers than in the previous variant. Large coefficient va-

6 278 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Table 1. Sub-association Hacquetio-Fraxinetum excelsioris typicum in the Cieszyńskie Foothills and adjacent areas Association Hacquetio-Fraxinetum excelsioris Marinček 1990 Subassociation Variant typicum brzostowy z Ulmus scabra with Ulmus scabra Successive No of record Terrain No of record Location Górki W. Date - day Date - month Year layer Exposure NW SW SW WNW W SW WSW SW WSW W SSW W Slope degree Density of tree layer a Density of tree layer a Cover of shrub layer b Cover of herb layer c Cover of moss layer d Area of record [m 2 ] No. Number of species Trees: V Fraxinus excelsior a a b c I Acer pseudoplatanus a a b c VI Acer campestre a b c I Ulmus scabra a a b c I Tilia platyphyllos a b c

7 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Hacquetio-Fraxinetum excelsioris Marinček 1990 typicum poor (vegetation quantity) disturbed Constancy Gułdowy Gułdowy Gułdowy Gułdowy Gułdowy Gułdowy Gułdowy Żywiec Ogrodzona Dzięgielów Dzięgielów Dzięgielów Dzięgielów Dzięgielów Dzięgielów Dzięgielów Dzięgielów Dzięgielów SW SW SW SW WSW W ESE NNW PLS W W NW NW W N N N PLS V III V V IV V III IV II I. I IV. I II I.. I

8 280 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): III Carpinus betulus a a b c III Tilia cordata a a b c IV Fagus sylvatica a a b III Cerasus avium a a c VI Acer platanoides a b c Quercus robur a a b c Alnus glutinosa a Alnus incana a c ii Padus avium s.s. a a b c Picea abies a Larix decidua decidua a Betula pendula a Sorbus aucuparia s.s. b Shrubs: Sambucus nigra b c VI Corylus avellana b c Crataegus laevigata b c VI Lonicera xylosteum b c V Hedera helix b c

9 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): V IV III IV V IV V V I. II I II. II I IV.. II V V II IV I I III II I II I III I II II III III II III II III I I. I I V IV III IV IV III. III II IV II III I IV III III II II II II

10 282 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Crataegus monogyna b c VI Euonymus europaeus b c VI Viburnum opulus b c V Ribes uva-crispa b V Daphne mezereum b Herbs: Ch. et *Dif Hacquetio-Fraxinetum excelsioris Marinček I 1990 et **Tilio platyphyllis-acerenion pseudoplatani Klika 1955 Hacquetia epipactis *Symphytum tuberosum *Ranunculus lanuginosus **Actaea spicata Fragaria moschata *Arum alpinum II Ch. Alno-Ulmion minorisbr.-bl et R. Tx Ficaria verna Anthriscus nitida Circaea lutetiana Stachys sylvatica III Ch. Carpinion betuli Issler 1931 em. Oberd.1953 Lathyrus vernus Vinca minor Galium schultesii IV Ch. Fagion sylvaticae R.Tx.et Diem Galium odoratum Dentaria glandulosa V Ch. Fagetalia sylvaticae Pawł Asarum europaeum Galeobdolon luteum s.l Mercurialis perennis Primula elatior Carex sylvatica Polygonatum multiflorum R Pulmonaria obscura Hepatica nobilis Isopyrum thalictroides Euphorbia amygdaloides Glechoma hederacea Paris quadrifolia Allium ursinum Euphorbia dulcis

11 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): II. III II II II. II I.. I I I I I I R V V V V V V. IV V IV II IV..... R I I. I II.. I II.. I III V III IV III II I II.. I I I I I II I II IV III I I I. I II V II I. III II V V IV V V V III IV II V V IV III V IV IV V III III IV R..... IV V I III I V IV III III V III I III II II I I III II I III I II R III I I II.. I III.. I

12 284 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Ajuga reptans Impatiens noli-tangere Lilium martagon Adoxa moschatellina Sanicula europaea Corydalis cava Gagea lutea VI Ch. Querco-Fagetea Br.-Bl. et Vlieg Anemone nemorosa Aegopodium podagraria Geum urbanum Milium effusum Viola reichenbachiana Brachypodium sylvaticum Rubus hirtus Melica nutans Catharinea undulata VII Ch. Betulo-Adenostyletea Br.-Bl Petasites albus VIII Other: Chaerophyllum temulum Oxalis acetosella Urtica dioica Galium aparine s.s Alliaria petiolata Lamium album Brachythecium sp Eurhynchium sp Occasionally species: Aesculus hippocastanum 204:R; Athyrium filix-femina 213:R; Caltha palustris 5:1; Carex digitata 107:+; Chrysosplenium alternifolium 12:+; Corydalis solida 109:+; Dentaria bulbifera 102:1; Dryopteris filix-mas 107:+; Galanthus nivalis 12:1; Geranium phaeum 105:R; Hypericum hirsutum 214:+; Impatiens parviflora 303:+; Lamium maculatum 26:+; Luzula luzuloides 5:+; Maianthemum bifolium 5:+; Marchantia polymorpha 5:+; Mnium undulatum 12:+; Plagiothecium laetum 205:+; Plagiotecium sp. 303:+; Poa nemoralis 205:1; Quercus rubra 215:+; Rubus caesius 303:+; Rubus idaeus 214:1; Rosa canina (b) 23:R; Salvia glutinosa 214:+; Senecio fuchsii 214:+; Stellaria holostea 105:R; Viola mirabilis 206:+. Ch. characteristic species; Dif* local differential species lues of coverage were attained for Mercurialis perennis, Symphytum tuberosum, Ficaria verna, Anemone nemorosa, Polygonatum multiflorum and Pulmonaria obscura, which was not noted in the previous variant The disturbed variant, like the poor variant, is characterised by an unnatural, single-layered tree stand, which consists of Fraxinus excelsior, Larix decidua, Picea abies, as well as a smaller numbers of Alnus glutinosa and Betula pendula, mostly from plantings. The only substantial species in the shrub layer is Sambucus nigra. Hacquetia epipactis is the only plant characteristic of the association. The forest floor is generally very poor, with only Fagion sylvaticae being distinguished with a larger coverage ratio compared to other variants, and this is due to two species Galium odoratum and Dentaria glandulosa. Also important are Mercurialis perennis, Hepatica nobilis and Anemone nemorosa The Hacquetio-Fraxinetum excelsioris allietosum sub -association (Table 2) This sub-association is distinguished primarily by the mass occurrence of Allium ursinum. Despite this, the phytocoenosis included in this syntaxon is characterized by a large number of species, more than in the typicum sub-association (by an average of three in specific surveys). The tree stand is made up of

13 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): II I I I II. I II I... R I II. I II I II. I II. I IV IV V V II IV V III V II II III II III II II IV. II II III.. I I I II I I.. I I I. I II I II I I II I III I I II I I II II. I I II I I III II I II II III I II multiple species and consists of two layers. The emergent layer most often consists of Fraxinus excelsior, Acer pseudoplatanus, Fagus sylvatica, Tilia cordata and Ulmus scabra; while the second tree layer usually has Carpinus betulus, and more rarely, Tilia cordata. The only unnatural element in the tree layer of this sub-association appears to be Quercus robur. The shrub layer has a very large proportion of Sambucus nigra, consisting of over 50% of coverage, and more rarely Crataegus monogyna. The herb layer is characterised by % coverage, with the aforementioned Allium ursinum, as well as considerable proportions of Mercurialis perennis, Asarum europeum, Ficaria verna and Galeobdolon luteum. Hacquetia epipactis has a low abundance, small clumps and long stems. Two variants were distinguished in the allietosum subassociation typical and with Fagus sylvatica: The typical variant is characterised by a significant share of Fraxinus excelsior and Quercus robur in the upper, emergent layer, while Carpinus betulus, Acer campestre and Tilia cordata are the most abundant in the second, tree layer. The shrub layer is represented by Sambucus nigra and Crataegus monogyna as well as C. laevigata. The forest floor has a much larger share of individual species than in the Fagus sylvatica variant. Hacquetia epipactis is characterised by small numbers, in contrast to the large proportion of species from the Alno-Ulmion alliance, especially Ficaria verna; as well as from the Fagetalia sylvaticae order, in which Mercurialis perennis is distinguished. Aegopodium podagraria, from the Querco-Fagetea class, has a large coverage ratio. It seems that if it were not for the large proportion of Quercus robur, this variant could be considered completely natural and optimal for Hacquetia epipactis, subject to interspecific competition The with Fagus sylvatica variant in the tree layer is also characterized by the presence of Tilia cordata, Acer pseudoplatanus, Quercus robur and Larix decidua in addition to the species from which it derives its name. The shrub layer is actually represented by only Sambucus nigra. As in the pre-

14 286 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Table 2. Sub-association Hacquetio-Fraxinetum excelsioris allietosum in the Cieszyńskie Foothills and adjacent areas Association Hacquetio-Fraxinetum excelsioris Marinček 1990 Subassociation allietosum Variant typical Successive No of record Terrain No of record Location Wiślica Wiślica Ochaby Ochaby Wiślica Wiślica Wiślica Wiślica Wiślica Gułdowy Goleszów Goleszów Goleszów Date - day Date - month Year Exposure warstwa / layer E ESE E NE ESE E E NE NE W WSW SW SW Slope degree Density of tree layer a Density of tree layer a Cover of shrub layer b Cover of herb layer c Cover of moss layer d Area of record [m 2 ] No. Number of species Trees: VI Fraxinus excelsior a a b c II Acer pseudoplatanus a b c VII Acer campestre a a b c II Ulmus scabra a a b c II Tilia platyphyllos b c

15 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): typical Hacquetio-Fraxinetum excelsioris Marinček 1990 allietosum bukowy z Fagus sylvatica with Fagus sylvatica Bażanowice Constancy Bażanowice Bażanowice Wilamowice Wilamowice Wiślica Wiślica Wiślica Leszna G. Rozumice S W NNE NW WSW W NW ENE NE SSW S NW S SSE SSE N NNW SE SW SE E R R R V III V II V III III I III II III II I I I

16 288 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): IV Carpinus betulus a a b c IV Tilia cordata a a b c V Fagus sylvatica a a b c IV Cerasus avium a VII Acer platanoides a Quercus robur a a Alnus glutinosa a Abies alba a III Padus avium s.s. b c Picea abies a a b Larix decidua decidua a III Ulmus laevis a b III Ulmus minor a Betula pendula a a Sorbus aucuparia s.s. c Shrubs: IX Sambucus nigra b c VI Hedera helix b c Crataegus monogyna b c VII Lonicera xylosteum b c VII Euonymus europaeus b c VII Corylus avellana b c

17 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): V V V IV V V II V III I. I +.. I. I III IV III I. I.. R. I I I. I II II II I III II I I... I. I I I I I. I R R R V V V II III III I II II II I II II II II I II II

18 290 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Crataegus laevigata b c VII Viburnum opulus c VI Daphne mezereum c VI Cornus sanguinea b Herbs: I Ch. et *Dif Hacquetio-Fraxinetum excelsioris Marinček 1990 allietosum *Allium ursinum Hacquetia epipactis R *Ranunculus lanuginosus *Symphytum tuberosum Fragaria moschata Arum alpinum II Ch. et *Dif Tilio platyphyllis-acerenion pseudoplatani Klika 1955 *Actaea spicata *Veronica montana III Ch. Alno-Ulmion minoris Br.-Bl et R. Tx Ficaria verna Anthriscus nitida Circaea lutetiana Stachys sylvatica Lamium maculatum Chrysosplenium alternifolium Geranium phaeum Festuca gigantea IV Ch. Carpinion betuli Issler 1931 em. Oberd.1953 Lathyrus vernus Vinca minor Lathrea squamaria s.s Galium schultesii V Ch. Fagion sylvaticae R.Tx.et Diem Galium odoratum Dentaria glandulosa VI Ch. Fagetalia sylvaticae Pawł Mercurialis perennis Asarum europaeum Galeobdolon luteum s.l Pulmonaria obscura Primula elatior R Glechoma hederacea Polygonatum multiflorum Hepatica nobilis Euphorbia amygdaloides Impatiens noli-tangere

19 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): II I II..... R I. I.... R R R I II I... I. I V V V R V V V IV III III III V III. R. II. I I I. + R II I II II I I III III III III I III III III III III II II R. + III II II I. I R R I I I I. I + R II III II.. 1 I. I I I I I I I III III III I I I V V V IV V V V IV V R.. IV II III IV II III R IV I III III V III II III III II III III II I II

20 292 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Paris quadrifolia Isopyrum thalictroides Euphorbia dulcis Ajuga reptans Carex sylvatica Adoxa moschatellina Lilium martagon Galanthus nivalis Sanicula europaea Corydalis cava Dryopteris filix-mas Astrantia major VII Ch. Querco-Fagetea Br.-Bl. et Vlieg Anemone nemorosa Aegopodium podagraria Milium effusum Viola reichenbachiana Geum urbanum Geranium robertianum Brachypodium sylvaticum Poa nemoralis Salvia glutinosa Campanula trachelium Melica nutans VIII Ch. Betulo-Adenostyletea Br.-Bl Petasites albus Senecio nemorensis s.s IX Ch. Epilobietea angustifolii R. Tx. Et Prsg Fragaria vesca Senecio fuchsii X Inne / Other: Urtica dioica Oxalis acetosella Galium aparine s.s Maianthemum bifolium Alliaria petiolata Athyrium filix-femina Impatiens parviflora Dryopteris carthusiana Hypericum hirsutum Chaerophyllum hirsutum s.s Carex brizoides Occasionally species: Anemone ranunculoides 526:+; Caltha palustris 5:1; Carex panicea 617:+; Carex pilosa 34:3; Carex sp. 540:+; Colchicum autumnale 552:1; 526:R; Deschampsia caespitosa 415:+; Dryopteris dilatata 414:+; Equisetum arvense 526:+; 617:+; Eurhynchium sp. 314:+; Fissidens taxifolius 512:+; Melandrium rubrum 540:+; Myosoton aquaticum 526:+; Phyteuma spicatum s.s. 414:R; Plagiotecium sp. 303:+; Pteridium aquilinum 540:+; Pyrus communis (b) 526:+; Rubus hirtus 616:+; Rubus idaeus 214:1; Rubus plicatus 616:+; Rumex sp. 540:R; Rumex sanguineus 616:+; Scilla bifolia 526:+; Scrophularia nodosa 526:R; Stellaria holostea 540:+; Taraxacum sect. vulgaria 552:R; 525:+ Ch. characteristic species; Dif* local differential species

21 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): II II II II I II III II II II. II II II II II. II I I I I I I R + I II I II. I. R I I I I. I IV V IV IV IV IV III III III R. II II II III I II II. II I I I I I I I I I R R I. I I I I. I I. I R R II I I I I I IV I III II II II III I II R. I II I I. I I I I II. I I I I I. I I. I I. I vious variant, the forest floor is dominated by Allium ursinum, with Hacquetia epipactis characterised by a larger share, and Ficaria verna occurring sporadically in contrast to the previous variant. Significantly greater consistency is achieved by Galium odoratum, Mercurialis perennis, Asarum europaeum and Anemone nemorosa. The remaining species are more common in the typical variant, as is the average number of total species in individual records (by an average of five species).

22 294 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Carpinion betuli 2.1. Tilio cordatae-carpinetum betuli, fragments of the association In contrast to the Hacquetio-Fraxinetum excelsioris, the phytocoenosis of Tilio cordatae-carpinion betuli with Hacquetia epipactis has this plant most often found in patches at isolated sites. Only one Tilio cordatae-carpinion betuli was noted in the Cieszyńskie Foothills (phytosociological record no. 11). All recorded phytosociological sites classified to this association are distinguished by flat or slightly inclined surfaces. The tree layer usually consists of Quercus robur, Acer pseudoplatanus and Carpinus betulus, which occur in both tree layers. Insignificant proportions of Larix decidua and Alnus glutinosa are also found here. The shrub layer consists of Sambucus nigra, Corylus avellana and Viburnum opulus. Hacquetia epipactis has an average coverage ratio, approximately at an intermediate level with Ulmus scabra variant and the disturbed variant of Hacquetio-Fraxinetum excelsioris typicum. The herb layer lacks the species characteristic for the Fagion sylvaticae alliance, as well as for the Carpinion betuli alliance, including those characteristic for the association. Significant coverage is provided by Polygonatum multiflorum, Hedera helix, Pulmonaria obscura, Asarum europaeum, Aegopodium podagraria and Galeobdolon luteum. The proportions of Paris quadrifolia, Ficaria verna and Oxalis acetosella are also quite significant Galio sylvatici-carpinetum betuli, fragments of the association This association was only identified in Grudzice. Fragments of this association have very few species, although the characteristic Ranunculus auricomus occurs here. The tree stand has a two-layered structure. The emergent layer consists of Quercus robur, Tilia cordata and Ulmus laevis, while the canopy layer is made up of Carpinus betulus and Acer pseudoplatanus. The shrub layer is very dense and occupies 50% of the surface. It is made up of Sambucus nigra, Acer pseudoplatanus and A. platanoides, Fraxinus excelsior, Tilia cordata and Padus avium. Hacquetia epipactis occupies very small fragments of the forest floor. In addition, the forest floor is very poor in species and contains small numbers of Aegopodium podagraria, Anemone nemorosa, Ficaria verna, Viola reichenbachiana, Ribes uvacrispa and many seedlings of the main tree species. A significant proportion of Hedera helix also occurs. 4. Discussion The high interest in Hacquetia epipactis over many years has generated various publications written by authors representing diverse fields of science, from different backgrounds and periods, resulting in much contradictory and not always reliable information, which cannot be unequivocally confirmed. The earliest studies noted the close association of Hacquetia epipactis to mixed deciduous forests. Hegi (1926) reports that in Europe, Hacquetia epipactis occurs in deciduous forests, as well as in alder forests and high in the mountains on grassy slopes adjacent to dwarf mountain pine. Then, these forests were included into the Fagetalia order (Koczwara 1960). Studies from Poland mention H. epipactis as belonging to a specific association. Kozłowska (1936) included H. epipactis in the Fagetum calcareum cieszynicum association, but the literature most often reports that its optimal occurrence is in hornbeam associations (Tumidajowicz 1964; Pelc 1969; Guzikowa 1970; Duda et al. 2001; Henel 2006; Gajewski et al. 2011). Wika et al. (2014) described H. epipactis in a fertile beech forest as a variant with Hacquetia epipactis (a site in the Wiślicka Escarpment reserve). Henel (2006) also lists the riverine ash-alder and elm-ash and mountain alder marshes as places where H. epipactis occurs. In Slovakia in turn, this plant was listed in the Vápnitá Lipová Javorina vyššieho stupňa assemblage, which belongs to the Tilieto-Aceretum alliance (Hančinský 1972). In 1990, Marinček described the limestone association Hacquetio-Fraxinetum excelsioris from the Alps, and then identified it in Austria (Mucina et al. 1993). Its centre of distribution is located in the Julian Alps, Austria and Croatia. In Poland, the description of this syntaxon is from the Wiślicka Escarpment, Municipal Woods on the Puńcówka Brook and Municipal Woods on the Olza River reserves, while fragments have been found in the Municipal Woods of Błogocice nature landscape park complex (Różański 2001a, 2001b; Różański, Dziedzic 2003). The mosaic of habitats in the Cieszyńskie Foothills and selected sites from all over Poland were not easy to analyse and difficulties were exacerbated by the slight differences in the values of similarities between groups of records and individual ones, which represented very similar habitats serving as the biochore for one plant species Hacquetia epipactis. During the analysis, it was found that the fertile deciduous forests with H. epipactis located in the Cieszyńskie foothills, classified earlier as lowland hornbeam or alluvial forests, can be identified as the Hacquetio-Fraxinetum excelsioris association of the Tilio- Acerion alliance (Różański 2001a, 2001b; Różański, Dziedzic 2003), and only a few phytocenoses with sites of Hacquetia epipactis represent non-typical hornbeam communities of the Carpinion betuli alliance. Additionally, the research conducted shows that the optimum occurrence of Hacquetia epipactis in a compact range in Poland is the Hacquetio-Fraxinetum excelsioris association, despite the fact that in many places this association is strongly altered or difficult to identify, and additionally lacking those species that are characteristic of the southern European range. This problem was the basis for distinguishing two, differing sub-associations (typicum and allietosum). Marinček (1990) distinguished two sub-associations dentarietosum trifoliae (including the variant with Allium ursinum) and omphalodetosum. None of the species distinguishing the

23 M. Mijal / Leśne Prace Badawcze, 2015, Vol. 76 (3): Slovenian sub-associations occurs in the phytocenoses with H. epipactis in Poland. The classification of phytosociological records showed that as much as 78% of the total area belongs to Hacquetio-Fraxinetum excelsioris, 12% represents hornbeam associations and 10% are unidentified phytocenoses. The characteristic and differential species of the Hacquetio -Fraxinetum excelsioris association, such as Lamium orvala, Helleborus odorus, Oryzopsis virescens, Crocus neapolitanus, Omphalodes verna often have a southern European range and do not exist in Poland. Others, such as Actaea spicata, Orchis pallens and Fragaria moschata are very rare in the Hacquetio-Fraxinetum excelsioris (Tables 1 and 2). This may be because Hacquetio-Fraxinetum excelsioris is very poorly known in Poland and is represented by a very impoverished form of this syntaxon at the extreme edge of its range. The dominant and differential association species most often occurring with H.epipactis the association, i.e. Symphytum tuberosum, Allium ursinum, Ranunculus lanuginosus, Corydalis cava, Polygonatum multiflorum and Isopyrum thalictroides were also listed by Mucina et al. (1993) from Austria. Hacquetia epipactis occupies specific microhabitats and does not tend to spread. Tumidajowicz (1964) noted the absence of H. epipactis in adjacent forests with similar habitat conditions, which is characteristic of the Mogilany site; Gajewski et al. (2011) confirmed that this species has not spread there. The same is true for the Cieszyńskie Foothills, where the plant remains in specific locations, does not change its site nor colonise new ones, and only exhibits annual fluctuations in the size of its population (Różański 2001b; Różański, Dziedzic 2003). Protecting the phytocoenoses of the Hacquetio-Fraxinetum excelsioris will enable the maintenance and survival of Hacquetia epipactis as a species associated with this syntaxon. Four locations of H. epipactis in the Cieszyńskie Foothills are in protected nature reserves. In addition, foresters in woods administered by the State Forests ensure the special protection of this species at sites that are outside of statutorily protected areas. Rational forest management, due to the need to maintain the phytocoenoses occupied by this plant, can contribute to its effective active protection, which has been signalled in various studies of H. epipactis. The most important protective actions include not allowing the excessive natural regeneration of species unrelated to the Hacquetio-Fraxinetum excelsioris community and avoiding excessive shade and sunlight (Duda et al. 2001; Malara et al. ; Henel 2006). Some believe that the fertile soils of the Cieszyńskie Foothills, which today are mostly crop fields or abandoned agricultural lands, indicate that they were covered by primary forests in the past (Wilczek et al. 2014), probably with a much higher number of patches of the Hacquetio-Fraxinetum excelsioris community than today. Evidence of this may be the occurrence of Hacquetia epipactis in each, even small forested site of this area, although in limited and specific micro-habitats. 5. Summary and conclusions Based on the study, the following conclusions have been formulated: 1. The main assemblage that includes Hacquetia epipactis in Poland is Hacquetio-Fraxinetum excelsioris, recorded in nearly all the sites analysed in the Cieszyńskie Foothills, as well as outside of it (Żywiecka Valley, Silesian Foothills and fragmentarily in Silesian Opole). 2. The assemblages of isolated sites of Hacquetia epipactis are represented by two hornbeam associations fragments of Tilio cordatae Carpinetum betuli (Wieliczka Foothills, the Upper Warta Depression) and fragments of Galio sylvatici Carpinetum (Grudzice). 3. Some of the phytocoenoses are very strongly distorted, making it impossible to classify them (the Silesian Beskid and Zywiecki Beskid). Other phytocoenoses are transitional and were not classified because they diverged from the diagram. 4. The sub-associations typicum and allietosum and the variants for each of them were distinguished in the Hacquetio -Fraxinetum excelsioris association for the first time in Poland. 5. The following species, which most frequently occur with Hacquetia epipactis, should be acknowledged as distinguishing the association: Symphytum tuberosum, Allium ursinum, Ranunculus lanuginosus and Corydalis cava. Conflict of interest and financial support The author declares no potential conflicts of interest. Acknowledgements My sincere thanks to Dr. Eng. Wojciech Różański and the reviewers for their help, useful counsel and valuable advice. References Duda J., Puchalski J., Szendera W Badania nad rozmieszczeniem i generatywnym rozmnażaniem cieszynianki wiosennej Hacquetia epipactis (Scop.) DC. Biuletyn Ogrodów Botanicznych 10: Dzwonko Z. 1977a. The use of numerical classification in phytosociology - Zastosowanie klasyfikacji numerycznej w fitosocjologii. Fragmenta Floristica et Geobotanica, series Polonica 23(3-4): Dzwonko Z. 1977b. Numeryczna klasyfikacja zbiorowisk leśnych Gór Stołowych (Polskie Karpaty Wschodnie). Fragmenta Floristica et Geobotanica, series Polonica 23(3-4): Dzwonko Z., Grodzińska K Numerical classification of epilitic and xerothermic communities in the Pieniny Mountains (Western Carpathies). Fragmenta Floristica et Geobotanica, series Polonica 25(4):